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Transmission of Grapevine Viroids Is Not Likely to Occur Mechanically by Normal Pruning

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Transmission of Grapevine Viroids Is Not Likely to Occur Mechanically by Normal Pruning Vitis 34 (2), 119-123 (1995) Transmission of grapevine viroids is not likely to occur mechanically by normal pruning by 1 U. STAUB, H. PouvKA, J.V. HERRMANN ) and H.J. GRoss Bayerische Julius-Maximilians-Universitat, Institut fiir Biochemie, Biozentrum,Wiirzburg, Germany 1 ) Bayerische Landesanstalt fiir Weinbau und Gartenbau, Veitshochheim, Germany S u m m a r y : In epidemiological studies the viroid distribution in two local vineyards was determined. Grapevine leaves of different varieties were collected, total RNA isolated and viroid detected by northern blot analysis and/or reverse transcription followed by PCR amplification. Nearly each sample was infected with the grapevine variant of Hop Stunt Viroid and approximately each second additionally with Grapevine Yellow Speckle Viroid l. Grapevine Yellow Speckle Viroid 2, a third grapevine viroid, was not found. Both grapevine viroids occurred in chlorotic plants as well as in plants without symptoms. In order to investigate viroid spreading through mechanical transmission accomplished during routine cultural practices, the distribution patterns in the two vineyards were analysed. Our results indicate that grapevine viroids are mainly propagated through systemic transmission upon grafting. The examination of different rootstock clones from Northern Italy, which are used for grafting in Germany, further demon­ strates that in this case propagation is not due to rootstocks containing viroid but is more likely to occur via infected scion varieties. K e y w o r d s : grapevine viroids, northern blot analysis, RT/PCR, rootstock, vine variety. Introduction has been isolated from Vitis vinifera cv. Riesling (PucHTA et al. 1988) and the rootstock 5BB (PucHTA et al. 1989). Viroids are the smallest known pathogens infecting only We performed this study with the aim to determine higher plants (RIESNER and GRoss 1985). They exist as cir­ the viroid distribution in two local vineyards. The analy­ cular, single-stranded RNA molecules with a length of 240- sis of viroid distribution patterns within a vineyard and in 600 nucleotides. Viroids of grapevine are distributed world­ different rootstock clones from Caprino Veronese (North­ wide and have been detected and characterized in Australia em Italy), which are routinely used for grafting in Ger­ (KoLTUNow andREzAIAN 1988 and 1989; REZAIAN et al. 1988; many, allows an insight into the mechanisms of viroid REZAIAN 1990), Germany (PUCHTA et al. 1988 and 1989), spreading. Japan (SANO et al. 1985), California (SzYcHowsKI et al. 1988) and Spain (GARCiA-ARENAL et al. 1987). From the five Materials and methods known grapevine viroids HSVdg (Hop Stunt Viroid grape­ vine), GYSVd1 (Grapevine Yellow Speckle Viroid 1), P 1 an t s : In a local vineyard (vineyard A) young GYSVd2 (Grapevine Yellow Speckle Viroid 2) that was leaves from Vitis vinifera cvs Bacchus, M tiller-Thurgau formerly named GV1b, CEVdg (Citrus Exocortis Viroid (rootstock 5 C), Kemer and Silvaner (rootstock SO 4) were grapevine) and AGVd (Australian Grapevine Viroid), only collected and stored at -20 oc until use. During the hot GYSVd1 and GYSVd2 were reported to induce symptoms summer of 1993 plants of cvs Bacchus and Miiller-Thurgau after infection ofviroid-free grapevine seedlings (KoLTUNOW were severely chlorotic whereas plants of cvs Kemer and et al. 1989). Grapevine viroids belong to the two main viroid Silvaner expressed no symptoms. For comparison we col­ groups that can clearly be distinguished from each other lected leaves of cvs M tiller-Thurgau and Kerner on by differences in their central conserved region (CCR) rootstock SO 4 in another local vineyard (vineyard B). (KEESE and SYMONS 1985). HSVdg is a member of the Even in the hot summer 1993 these plants expressed no PSTVd (Potato Spindle Tuber Viroid)-group and GYSVd1, chlorotic symptoms. In order to elucidate whether viroid GYSVd2 and AGVd are members of the ASSVd (Apple infection and spreading derives from the rootstock, leaves Scar Skin Viroid)-group (HASHIMOTO and KoGANEZAWA from different clones in Caprino Veronese (Northern Italy) 1987). In comparative studies between Europe and Cali­ were also obtained and assayed for viroid infection. fornia (SzYCHOWSKI et al. 1991) and California and Aus­ tralia (REZAIAN et al. 1992) it turned out that nearly all sam­ E x t r a c t i o n o f n u c 1 e i c a c i d s : Grapevine ples studied were infected with the latent and symptomless leaves (0.25 g samples) were ground in liquid nitrogen HSVdg (SANO et al. 1985). Furthermore, many cultivars with a mortar and a pestle. The powder was transferred were coinfected with either GYSVd1 or GYSVd2, or with into a 2.0 ml microcentrifuge tube and 800 111 homogeni­ both. Plants solely infected with GYSVd1 or GYSVd2 with­ zation buffer (0.2 M boric acid, 10 mM Na2EDTA, pH out a HSVdg-infection were not found. In Germany HSVdg adjusted to 7.6 with Tris), 16 111 25% SOS and 16 111 of Correspondence to: Prof. Dr. H. J. GRoss, Universitat Wiirzburg, Institut fiir Biochemie, Biozentrum, Am Hub land, D-97074 Wiirzburg, Germany 120 U. STAUB, H. PoLIVKA, J.V. HERRMANN and H.J. GRoss 2-mercaptoethanol were added. The mixture was shaken nucleotide primers, were employed for RT/PCR (STAUB et for 10 min with 800 ~1 phenol/chloroform/isoamyl alco­ al. 1995). Amplification products were separated in a 2.0 % hol (25/24/1, v/v/v) saturated with 10 mM Tris-HCl/1 mM agarose gel and visualized by ethidium bromide staining. Na2EDTA (pH 7.0) and centrifuged at 15000 g (4 °C) for 10 min. Phenol extraction was repeated and the aqueous N o r t h e r n b I o t a n a 1 y s i s f o r t h e d e­ phase finally extracted with 400 ~1 chloroforrn/isoamyl t e c t i o n o f g r a p e v i n e v i r o i d s : 5 jlg total alcohol (24/1, v/v). Nucleic acids were further purified ei­ RNA was separated in a 5% polyacrylamide/7 M urea gel ther by differential solvent precipitation (MANNING 1991) and electrotransferred onto a positively charged nylon or by an aqueous two-phase system (BEUTIIER et al. 1988) membrane (Qiabrane Nylon Plus; Qiagen, Hilden). Hybridi­ followed by DEAE-cellulose chromatography according zation was carried out at 60 °C with equal amounts of ra­ to STAUB et al. (1995). dioactively 5' end-labelled oligodeoxynucleotides HV1 and GV3 simultaneously (Tab. 1; STAUB et al. 1995). After Detection of viroids by RT/PCR: In hybridiziation the membrane was washed under stringent a coupled reaction (MYERS and GELFAND 1991) viroid-RNA conditions (0.1 x SSC, 0.1% SDS; 60 °C) and the result in 25 ng total RNA, isolated from grapevine leaves, was was documented by autoradiography. Signal intensities reverse transcribed (RT) and viroid-cDNAamplified (PCR). were quantified with a Molecular Dynamics Phosphor­ GV1 and GV2 (Tab. 1), two GYSVdl-specific oligodeoxy- Imager 425. Tab I e I Oligodeoxynucleotide primers used for RT/PCR studies and northern blot analyses name sequence specific for position within viroid domain the molecule GVl 5'-GCGGGGGTTC CGGGGATTGC-3' GYSVdl 341 - 360 TL GV2 5'-taagaggtct ccggatcttc ttgc-3' GYSVdl 361- 17 TL GV3 5'-ACCCCTICGT CGACGACG-3' GYSVdl 98- 115 CCR GYSVd2 HVl 5'-GTIGCCCCGG GGCTCCTI-3' HSVdg 71 - 88 CCR Sequences in capital letters are complementary to the corresponding viroid, the sequence of GV2 (in lower case letters) is the DNA homologue of the viroid sequence. GYSVdl: Grapevine Yellow Speckle Viroid 1; GYSVd2: Grapevine Yellow Speckle Viroid 2; HSV dg: Hop Stunt Viroid grapevine; T L: left terminal region of the molecule and CCR: central conserved region of the viroid according to KEESE and SYMONS (1985). Results Samples of other varieties in vineyard A were exam­ ined in the same way. The results are summarized in Fig. 3. Viroid distribution in two local Nearly each plant is infected with HSVdg and about 50% v i n e y a r d s : In the investigated areas two of the five of the samples studied are additionally infected with known grapevine viroids, HSVdg and GYSVd1, were iden­ GYSVdl. 81% of chlorotic cv. Bacchus (rootstock 5 C) tified by RT/PCR and northern blot analyses. For the de­ samples and 46% of chlorotic cv. Muller-Thurgau tection of GYSVd1 in different samples RT/PCR was car­ (rootstock 5 C) samples, but only 29% of symptomless cv. ried out with the specific oligodeoxynucleotides GV1 and GV2. The expected PCR products with 367 bp length cor­ M 23456 7 responding to full-lenght GYSV dl were visualized by stain­ 4~9--.,.,._ ing with ethidium bromide (Fig. 1, lanes 3-7). 41~----­ In this epidemiological study grapevine viroids were .1 .'1- - 367bp 2-l~_../ mainly detected by northern blot analyses as shown in 1'){)__........ 147 ___........ Fig. 2. 5 ~g total RNA isolated from different samples of I 10 ___........ cv. Muller-Thurgau in vineyard A were fractionated by electrophoresis in a 5% polyacrylamide/7 M urea gel, Fig. I: Detection of GYSVdl by RT/PCR from young electrotransferred onto a positively charged nylon mem­ leaves of different grapevines. Lane M: marker pUC19/Hpa II fragments brane and hybridized with labelled oligodeoxynucleotides (length in bp). Lane 1: control; RT/PCR product from total RNA GV3 and HVl. Some of the assayed samples are viroid­ of a GYSVdl-infected plant (cv. Mtiller-Thurgau). Lane 2: con­ free (Fig. 2, lanes 4, 12) and others are infected with either trol; RT/PCR with ddHp instead of total RNA. Lanes 3-7: RT/ HSVdg (lanes 6, 9, 11, 13) or HSVdg and GYSVdl (lanes PCR products from total RNA of different samples of 3, 5, 7, 8, 10).
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